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FIELD GUIDE TO

SHARKS, RAYS & CHIMAERAS

OF EUROPE AND THE MEDITERRANEAN

Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540

In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TR press.princeton.edu

Princeton University Press is committed to the protection of copyright and the intellectual property our authors entrust to us. Copyright promotes the progress and integrity of knowledge. Thank you for supporting free speech and the global exchange of ideas by purchasing an authorized edition of this book. If you wish to reproduce or distribute any part of it in any form, please obtain permission.

Requests for permission to reproduce material from this work should be sent to Permissions, Princeton University Press

Copyright © 2021 by Princeton University Press

Illustrations copyright © 2021 by Marc Dando

Copyright in the photographs remains with the individual photographers. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the publishers.

British Library Cataloging-in-Publication Data is available

Library of Congress Control Number 2020944541

ISBN 978-0-691-20598-4

Ebook ISBN 978-0-691-21182-4

Production and design by WILDNATUREPRESS Ltd., Plymouth, UK

Printed in Italy

Cownose rays Rhinopteridae

Mobulid rays Mobulidae

● Electric rays Torpediniformes

Torpedo rays Torpedinidae

Torpedo ray species key guide

● Rhino rays Rhinopristiformes

Sawfishes Pristidae

Sawfish species key guide

Guitarfishes Rhinobatidae

Guitarfish species key guide

Giant guitarfishes Glaucostegidae

Comparison of shark-like rays and ray-like sharks 184

● SHARKS SELACHIMORPHA

Shark orders key guide

● Cow and frilled sharks Hexanchiformes 191

Cow sharks Hexanchidae 191

Frilled sharks Chlamydoselachidae 191

Cow and frilled shark species key guide 192

Cow and frilled shark species dentition guide 193

● Bramble sharks Echinorhiniformes/Echinorhinidae

● Dogfish sharks and related species Squaliformes

Dogfish sharks and related families key guide

Dogfish sharks Squalidae

Dogfish shark species key guide

Dogfish shark similar species comparison

Dogfish shark species dentition guide

Gulper sharks Centrophoridae

Gulper shark species key guide

Gulper shark species dentition guide

Lanternsharks Etmopteridae

Lanternshark species key guide

Lanternshark species dentition guide

Deepsea spined shark species comparison

Sleeper sharks Somniosidae

Sleeper shark species key guide

Sleeper shark species dentition guide

Roughsharks Oxynotidae

Roughshark species key guide and dentition guide

Kitefin sharks Dalatidae

Kitefin shark species dentition guide

Kitefin shark species key guide

● Angelsharks Squatiniformes/Squatinidae

Angelshark species key guide

Angelshark species dentition guide

● Carpetsharks Orectolobiformes

Nurse sharks Ginglymostomatidae

Whale sharks Rhincodontidae

Carpetshark species key guide

Carpetshark

Mackerel shark families

Sandtiger sharks Carchariidae and Odontaspididae

Sandtiger and goblin shark species dentition guide

Thresher sharks Alopiidae

Thresher

Basking

Mackerel

Large

Acknowledgements

A book like this can only come to fruition through the contributions of many individuals, either directly or indirectly. We are fortunate to work with amazingly talented and dedicated colleagues. There are far too many to name individually but we would particularly like to thank Mark Harris, FFC, Elasmobranch Studies, for his encyclopedic knowledge of all things tooth-related; Fabrizio Serena for his wide knowledge of this region; Ali Hood for shining a spot-light on shark conservation; Sarah Fowler, Cat Gordon, Rima Jabado, Peter Kyne, Matthias Stehmann, Guy Stevens and Simon Weigmann for numerous informative discussions and input.

We are extremely grateful for the excellent proofing skills of John Richardson and Peter Kyne; to Julie Dando for overseeing and managing this enormous project and to Robert Kirk for his belief and enthusiasm in this book.

Dave Ebert would like to thank Marsha Englebrecht and his parents, Earl and Margaret (Peggy) Ebert, for their support and encouragement.

Marc Dando especially thanks his family for their understanding, forbearance and support whilst working on this project, without their help and tolerance this guide would never have happened, so thank you, Julie, Ryan, Megan, Darren, Riley and Sonny.

NORTH ATLANTIC OCEAN

ARCTIC OCEAN

Regional seas covered in this book

This book is intended to be a guide to all known living species of sharks and their relatives, the skates, rays and chimaeras, found in the waters of the Northeast Atlantic and Mediterranean region. Seventy-nine species of shark, fifty-eight species of skate and ray (batoids) and nine species of chimaera are found in the region. These are variously distributed from coastal nearshore waters, including estuaries, out to depths of approximately 200m, along the outer continental shelf, and beyond that, hundreds to thousands of metres down the continental slope and beyond, where many fascinating and unusual looking deepsea species occur. While some species occurring in these waters have a global distribution, others

are endemic or near endemic to the Northeast Atlantic. The Mediterranean Sea also has several endemic species, as well as others that appear to have migrated from the Red Sea through the Suez Canal. Many of the species present in the Northeast Atlantic and Mediterranean region are well known, and often have a long history associated with fisheries, whereas others have only recently been documented as occurring here, including several species new to science.

We hope this guide will encourage and inspire more people to take an interest in sharks and their relatives, whether as a casual observer, or more actively through research, education, fisheries management or conservation.

European seas – biodiversity

Sharks and rays are among the most recognisable of fishes and, while most people easily recognise charismatic species such as the White Shark Carcharodon carcharias or Giant Manta Ray Mobula birostris, fewer are aware of the remarkable diversity of species that exist. Sharks come in a variety of sizes and shapes: from the Spined Pygmy Shark Squaliolous laticaudus that, with a maximum length of 22cm, you can hold in the palm of your hand; to the largest fish in the ocean, the Whale Shark Rhincodon typus, reaching 20m total length. However, only 18% of shark species have a maximum length exceeding two metres or a maximum weight exceeding 10kg. This is also true of rays and skates.

Sharks and their relatives, the skates, rays and chimaeras, collectively form the Class Chondrichthyes –a distinctive fish group also referred to as ‘cartilaginous fishes’ pertaining to their simple internal skeletons. In these fishes, cartilage replaces the more complicated internal calcified bony structures found in ‘bony fishes’. Globally, there are presently 540 known species of shark, 665 species of skate and ray (known collectively as batoids and affectionately as ‘flat sharks’) and 52 species of chimaera (or ‘ghost sharks’), an overall total of 1,257 species. In comparison, the far more diverse bony fishes total about 33,000 species.

The waters of the Northeast Atlantic and the Mediterranean region are home to nearly 12% of all known shark species, placing it among the world’s top 20 shark diversity hotspots, with a combined total of 146 species. In comparison, the top three global shark

ClassSubclassCohortOrder

Holocephali

Batoidea (batoids)

Chondrichthyes

Neoselachii

Selachii (sharks)

diversity hotspots are the regional waters of Australia (329 species), India (227 species) and Japan (212 species). Of the 146 European species, 137 occur in the Northeast Atlantic and just 86 in the Mediterranean. Seventy-seven species occur in both. The number is higher in the Northeast Atlantic because many deepwater species that live there in depths below 500m are missing from the Mediterranean, probably because the narrow, relatively shallow (300m) Strait of Gibraltar creates a barrier to their movement. There are about 23–25 Northeast Atlantic endemics (species that are found nowhere else in the world). The Mediterranean species list includes four sharks and five batoids that are not recorded in the Northeast Atlantic. These are mostly coastal warm water species. Two batoids are endemic to the Mediterranean, and three species have entered the Mediterranean from the Indian Ocean and Red Sea, through the Suez Canal. Their entry to the Mediterranean from the Red Sea is known as ‘Lessepsian migration’ (so named after Ferdinand de Lesseps, the engineer in charge of constructing the Suez Canal during the 1860s).

The known diversity of sharks and their relatives (hereafter collectively referred to as sharks) has increased exponentially over the past 19 years, with nearly 300 new species described between 2000–2018. This is over 20% of all shark species that have been described and named to date. However, in the 50 previous years between 19501999, only 330 species were described. The majority of recent discoveries have come from the Indo–Australian region, followed by southern Africa and the western

Chimaeriformes page 46

Rajiformes page 68

Myliobatiformes page 139

Torpediniformes page 166

Rhinopristiformes page 175

Hexanchiformes page 191

Echinorhiniformes page 198

Squaliformes page 200

Pristiophoriformes not in this region

Squatiniformes page 252

Heterodontiformes not in this region

Orectolobiformes page 258

Lamniformes page 273

Carcharhiniformes page 286

Table 1. The Class Chondrichthyes is divided into two major groups: the chimaeras, comprising the subclass Holocephali; and the sharks, skates and rays, forming the subclass Neoselachii. While the chimaeras all belong to a single order (Chimaeriformes), the sharks, skates and rays are separated into 13 orders: nine orders of shark-like fishes (cohort Selachii), and four of ray-like fishes (cohort Batoidea).

North Pacific. Although the majority of Northeast Atlantic and Mediterranean species were described in the 20th century, six new species have been named during the past 20 years, five were named from specimens collected in the region, while a sixth was described outside the region, but subsequently found to occur here.

Sharks are one of the most successful groups of fishes, having penetrated most of the world’s marine ecosystems, from tropical to high latitude waters over continental and insular shelves, out into the open ocean and the deepsea (>200m). Their success is not limited to marine ecosystems; in some regions certain shark and ray species either inhabit, or seasonally visit, estuarine and freshwater river ecosystems. The diversity of known species is greatest in tropical marine ecosystems, followed by the deepsea and temperate ecosystems, with oceanic and freshwater ecosystems having the lowest diversity. The majority of species (~55%) occur on continental shelves from the intertidal zone, including bays and estuaries, along coastal nearshore areas and out to about 200m depth. The diversity of these shelf species is greater in the tropics and lower in temperate seas. The remainder (~45%) are considered deepsea species. Species diversity in oceanic and freshwater habitats is comparatively low, making up only approximately 3% and 2%, respectively.

The modern taxonomic classification system for all organisms was designed by Carolus Linnaeus (1707–1778), a Swedish botanist, physician and zoologist. Linnaeus formalised the system for naming organisms, with each assigned a binomial (two-part) scientific name, usually based on Latin or Greek. The binomial scientific name consists of the genus (generic) and species (specific) name, both of which are written in italics. Therefore, when identifying or recording an organism, the binomial scientific name is given preference over its common name, which may vary from region to region.

The taxonomic system is hierarchal, with kingdom being the broadest or highest level of classification, followed in descending sequence by phylum, class, subclass, cohort, order, family, genus and species. In this book, the class Chondrichthyes is the highest taxonomic classification referred to, see Table 1.

The ordinal classification is relatively even between the Northeast Atlantic and Mediterranean, with 12 and 11 orders, respectively. The only order found in the Northeast Atlantic, but absent in the Mediterranean is the Orectolobiformes (carpet sharks). However, with each descending taxonomic level the difference between the two regions increases, see Table 2.

All the families in this book occur in the Northeast Atlantic. However, there are five families that are not represented in the Mediterranean, the Chlamydoselachidae (frilled sharks), Ginglymostomatidae (nurse sharks),

Table 2. Chondrichthyes diversity in the Northeast Atlantic and the Mediterranean regions, the greatest diversity being visible in the sharks.

Cohort/order

Chimaeriformes (chimaeras)

Batoidea (batoids)
Selachii (sharks)
Batoidea (batoids)
Selachii (sharks)

Rhincodontidae (whale sharks), Mitsukurinidae (goblin sharks) and Pseudotriakidae (false catsharks). In the batoids, two families are absent from the Mediterranean: the Arhychobatidae (softnose skates) and Pristidae (sawsharks). However, the Mediterranean is home to the genus Himantura, a genus not found in the Northeast Atlantic.

There are also several vagrant species, including two shark species: the Crocodile Shark Pseudocarcharhias

kamoharai and Night Shark Carcharhinus signatus

More recently, in 2015, a Sicklefin Chimaera Neoharriotta pinnata was discovered in the southern Bay of Biscay, extending its previous known range from Northwest Africa. Changes in oceanographic conditions, especially due to warming water temperatures, likely explains recent vagrant records of normally warm temperate to tropical species, while deepsea species may have been overlooked until recently.

European seas – habitats

Sharks, skates, rays and chimaeras can be found in a wide variety of different marine habitats. Some spend their whole life on the seabed (benthic species), whilst others cruise the high seas and never touch bottom (pelagic species). Many, including most chimaeras, lurk near the seabed where they hunt for bottom-living fish and invertebrates (bentho–pelagic species). Whether the seabed is rock, coral, sand, mud, or a mixture, there will be species living there, each adapted to its particular preferred habitat.

As well as these habitat and way of life considerations, sharks, skates, rays and chimaeras can be split into three ecological groupings, based on where they live in relation to continental landmasses and the seabed. Those species found from coastal nearshore waters to a depth of about 200m are referred to as continental shelf inhabitants.

Species inhabiting depths below 200m, and living on or near to the seafloor, are considered continental slope or deepsea species. Those species mostly living far off the bottom in the upper few hundred metres of the water column, and primarily away from continental landmasses, are referred to as oceanic species. This latter small group includes some species found around offshore islands and insular slopes, but not considered to be associated with continental landmasses. Species are further differentiated biogeographically according to the temperature of their home water, which might be tropical, temperate or polar. So, for example, a species living in the warm coastal waters off Italy would be a warm temperate continental shelf inhabitant, whereas those spending all of their time far out in the colder seas of the Northeast Atlantic are cold temperate oceanic species.

Map of Large Marine Ecosystems (LMEs) in our region

19 East Greenland Shelf

Barents Sea

Norwegian Sea

The continental shelf itself can be separated into two areas: a nearshore, or coastal habitat that includes bays and estuaries and which usually extends out to about 50m depth, and an offshore, or outer continental shelf habitat that extends from 50m to about 200m depth.

Many of the sharks, skates and rays that inhabit nearshore coastal waters will be familiar to anglers, divers and others who spend time near the sea. Chimaeras are unlikely to be encountered here, though there are some exceptions, such as the many steep-sided sea lochs found along the Norwegian coast, where some chimaeras may be found within diving depths.

Spotting sharks in offshore waters, including the outer continental shelf, pelagic and deepsea habitats, is a different prospect, but a variety of common oceanic sharks can be seen in pelagic waters not too far offshore, such as the Basking Shark Cetorhinus maximus or Porbeagle Shark Lamna nasus, but the only batoid that might be regularly encountered is the Pelagic Stingray Pteroplatytrygon violacea. Although the deepsea habitat is more diverse and supports a large number of species, the chance of encountering a deepsea shark, skate or chimaera is low, unless you are on a fishing boat or research ship.

Chondrichthyes species are can be seen in their habitat throughout the region, from angelsharks inhabiting shallow warm waters around the Canary Islands (left) to deepsea species like chimaeras in the cold waters off the Norwegian coast (right)

The European marine habitat is a vast area that encompasses the mostly cold-temperate Northeast Atlantic and the much warmer Mediterranean Sea. It comprises about 18 million square kilometres, an area nearly twice that of the European landmass, and has a coastline around 250,000km long. The region covered in this book also includes the islands of the Azores, Canaries and Madeira archipelagos. The Mediterranean Sea extends from the Strait of Gibraltar to the Bosphorus, and this book also includes the Black Sea.

The region includes two major FAO (Food and

Agriculture Organisation) fishing areas: the Northeast Atlantic (Major Fishing Area 27) and the Mediterranean and Black Seas (Major Fishing Area 37). It also encompasses 13 Large Marine Ecosystems (LMEs). LMEs are regions of the world’s oceans with distinct bathymetry, hydrography, marine productivity and food chain interactions. As their name suggests, these are large areas, in the order of 200,000km2 or greater, encompassing coastal waters, including river basins and estuaries, out to the edge of continental shelves and the outer margin of major ocean currents (see map opposite).

Table 4. Physical statistics of the North Atlantic, Mediterranean and Black Sea and the number of chondrichthyan species (sharks, rays, skates and chimaeras) known to occur in those areas

* 2 native, another 1 vagrant (Gymnura altavela) ** 2 native, another 4 vagrant (Squatina squatina, Alopias vulpinus, Scyliorhinus canicula and Sphyrna zygaena)

CONSERVATION AND MANAGEMENT

An overview

The evolution of conservation management activities in Northeast Atlantic and Mediterranean waters is complex, with multiple actors representing myriad interests. The European marine habitat is vast with a coastline of ~250,000km. This geographic scope presents a region with almost 40 coastal States complicated by extremes of wealth and political stability, of temperature and bathymetry. As well as comprising some of the most anthropogenically pressured habitats for elasmobranchs globally, it is also a region with sharks on the menu. In this chapter the term ‘sharks’ refers to all chondrichthyans (sharks, rays and chimaeras).

This section seeks not to present a comprehensive report on decades of conservation development and activity, but to deliver an overview, sign-posting readers to organisations and further materials, while profiling some of the species central to the conservation narrative of the region. The inauguration of the European Elasmobranch Association in 1996, and the closely associated inception of a number of national shark conservation organisations, provides a convenient 25 year timeframe (see below).

It is generally appreciated that sharks are inherently vulnerable to over-exploitation due to their late maturity, low fecundity and slow growth. The retention of sharks and batoids from target and mixed fisheries, and international market demand for shark and batoid products (including fins, meat and liver oil), has expanded over recent decades. Yet, where shark finning was previously the key driver for shark fisheries, the retention of sharks for meat is now an important and expanding consideration. Meat imports increased 42% by volume between 2000 and 2011. Pair this demand with sharks’ inherent biological vulnerability, and the result is declines in many elasmobranch populations. This is particularly apparent in Mediterranean and Northeast Atlantic waters, where over 53% and 32%, respectively, of chondrichthyans face an elevated risk of extinction, according to the IUCN Red List of Threatened Species.

NORTHEAST ATLANTIC AND

MEDITERRANEAN TIMELINE

This timeline illustrates conservation progress in the region over the past 25 years. It is not comprehensive, but serves to project the growing momentum of conservation management actions. Yet while listings increase, other urgent conservation priorities fail to secure much needed measures, and action and implementation often lag behind.

Fisheries interests in the region are significant. The long history of shark fisheries is evidenced by the number of gear types named after the sharks that they were designed to target, from the ‘squaenera’ deployed to target angelsharks (Squatina spp.) in Italy, to the ‘cazonal’ used to catch smoothhounds (Mustelus spp.) in the Canary Islands. Black and white archive photographs show fish quays laden with species now rarely encountered across vast tracts of their previous range. And, while coastal fisheries may now be depleted of previously common species, shark fishers continue to fill their holds from high seas waters.

Across the region, decades of poor catch reporting have left many species data-limited, with aggregated landings data (for example ‘sharks nei’ or ‘skates and rays nei’; ‘nei’ stands for not elsewhere included) often masking concerning changes in catch composition. Species-specific landings data are key to informed fisheries management, and improved species identification is central to this.

While the Mediterranean is characterised by small scale (mixed) fisheries, Northeast Atlantic shark fisheries are dominated by a small number of countries, accounting for a significant percentage of European Union (EU) and global shark landings. Spain ranks second in the world for shark catch, and has fleets active throughout the world’s oceans. Illegal, unreported and unregulated (IUU) fisheries are a focus of attention for many of the regional fisheries management organisations, such as International Commission for the Conservation of Atlantic Tunas (ICCAT) and the General Fisheries Commission for the Mediterranean (GFCM). (See Table 6 (p.26) for other regional bodies.)

The EU, with its 27 Member States and extensive fishing fleet, exerts both influence and impact throughout this region as a fishing power, and as the proponent for both regional and international regulation. The consequences of the UK’s exit from the EU in January 2020 are yet to be fully

1991–1994

Proposals to establish EEA discussed

1995

Barcelona Convention SPA/ BD signed EEA (see p.17 for country member groups established in this year)

1996

European Elasmobranch Association (EEA) established

appreciated. But, given the conservation stance favoured by the UK in recent decades, the loss of this key counterpoint to the more pro-sharkfishing Member States within the EU’s negotiating forum may be keenly felt.

Commercial fishers are not the only stakeholders with interest in sharks in the region. Recreational sea anglers have long targeted sharks for sport, albeit increasingly as catch and release. Ecotourism adds another facet to the burgeoning tourist industry, catering to an appetite for wildlife watching, while the coastal developments that cater to tourism may themselves damage vital nearshore habitats.

In 1999, the UN Food and Agriculture Organization (FAO) adopted the International Plan of Action for the Conservation of Sharks (IPOA-Sharks). In 2003, this approach was reflected in the publication of the voluntary Action Plan for the Conservation of Cartilaginous Fishes (chondrichthyans) in the Mediterranean Sea by the Barcelona Convention’s Regional Activity Centre for Specially Protected Areas (RAC/SPA). In 2009, the EU finally adopted the Community Plan of Action for Sharks (CPOASharks) – although both documents are yet to deliver on many of their objectives. In recent years, the development by NGOs of speciesspecific Action Plans has helped identify necessary actions at appropriate scales, and foster greater collaboration.

So, while recognising the troubling conservation status of the region’s elasmobranch populations, progress has been made within the 25 year timeframe presented here. Conservation and management of sharks has advanced at national, regional and international levels, although management for batoids has noticeably lagged behind that of sharks, and patchy implementation of voluntary measures and regulations hinders compliance. Key regulations and legislation are listed in Table 6 (p.26) and an overview of the region is presented over page.

1997

EEA (see p.17 for country member groups established in this year)

1998

Basking Shark UK protection

BASKING SHARKS Cetorhinus maximus (p.277)

Hunted for fins, meat and liver oil, Basking Sharks were caught commercially into the mid 1990s, eventually targeted solely for their high value fins.

Listed under UK domestic protection in 1998, EU Fishing Regulations in 2001, CITES Appendix II in 2003, and CMS Appendix I & II in 2005, Basking Sharks have successfully migrated from commercial targets to arguably one of the world’s most protected elasmobranchs. There are no legal directed fisheries in Northeast Atlantic or Mediterranean waters.

Much heralded as wildlife ambassadors, commercial interests in Basking Sharks now reflect the growing ecotourism sector. Similarly, citizen science initiatives embrace the public interest in reporting sightings, and submission of fin images can support photo­identification. On the water, codes of conduct for water­users help guide safe shark encounters.

Unfortunately incidents of bycatch, entanglement and vessel collision persist.

THE COMMON SKATE COMPLEX (pp.96–99)

Skate and ray nomenclature often causes confusion, with some species of the family Rajidae (the hardnose skates), being referred to as rays (e.g. Thornback Skate Raja clavata is often called the Thornback Ray). A diversity of colloquial common names further muddies the waters. And, with skate landed in aggregate in Northeast Atlantic waters until ~2008, changes in catch composition largely went unnoticed.

The large­bodied Common Skate Dipturus batis was historically a key part of the catch composition. Subject to overfishing, their numbers declined dramatically, seeing them listed as Critically Endangered (2006). Yet, the confusion here is not one of name alone – the Common Skate, no longer common. Genetic studies published in 2010 concluded that the Common Skate was not one, but two species. Some ninety years after two visually similar skate species had been conflated as Common Skate, they were separated once more. The two species identified as the Common Skate complex have new taxonomic names allocated: the Blue Skate Dipturus batis (p.96) and the more northerly, and larger, Flapper Skate Dipturus intermedius (p.98).

1999

First shark listed on the Convention on Migratory Species (CMS) Appendix II: Whale Shark (See Table 5); UN Food and Agriculture Organization (FAO) adopted the International Plan of Action for the Conservation of Sharks (IPOA­Sharks) First North Sea TAC for ‘skates and rays’

2001

Basking Shark first listed on Common Fisheries Policy

2002

First elasmobranchs listed on the Convention on International Trade in Endangered Species (CITES) Appendix II: Basking Shark and Whale Shark (See Table 5) SPA/BD ratified

2003

First EU Finning Regulation adopted; the Action Plan for the Conservation of Cartilaginous Fishes (Chondrichthyans) in the Mediterranean Sea adopted in the framework of the UNEP/MAP Barcelona Convention EEA (see p.17) contd. on p.16

The region

Chimaeras, batoids and sharks in the region 146

Chimaeras, batoids and sharks in the Mediterranean 86

Chimaeras, batoids and sharks in Europe (NE Atlantic) 137

CITES* Appendix I and II listed species in the region 18

CMS** Appendix I and II listed species in the region 23

*The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES)

**The Convention on the Conservation of Migratory Species of Wild Animals (CMS)

BLUE SHARKS AND MAKO SHARKS

Bycatch in high seas fisheries is a significant threat to pelagic elasmobranchs. Beyond the EEZ (Exclusive Economic Zone) of any one country, the high seas are regulated by Regional Fisheries Management Bodies.

Blue Shark Prionace glauca (p.343) is the most heavily fished species globally. Generally unregulated, in 2019 ICCAT adopted the first catch limit for sharks in international waters for Blue Sharks in the North and South Atlantic.

Shortfin Mako Isurus oxyrinchus (p.282), and Longfin Mako Isurus paucus (p.283), are exceptionally vulnerable and a pressing conservation concern. Repeatedly overlooked for management, Shortfin Mako is now subject to clear advice from ICCAT scientists, who in 2017 advised a prohibition on retention of mako in the North Atlantic. CITES Appendix II listing was adopted for Short and Longfin Mako in 2019.

IUCN Red List status

Analysis by the IUCN Species Survival Commission’s Shark Specialist Group estimates that, globally, 24% of chondrichthyan species are likely threatened with extinction. This high rate of risk, caused primarily by overfishing, distinguishes this group of fishes as among the most threatened of the world’s vertebrate groups. EU – % of species threatened

Europe Red List†

Critically Endangered (CR) 15

Endangered (EN) 17

Vulnerable (VU) 10

Near Threatened (NT) 14

Least Concern (LC) 48

Data Deficient (DD) 27 Total assessed 132

† EU Red List includes Mediterranean species †† Nieto et al. (2015)

2004

International Commission for the Conservation of Atlantic Tunas (ICCAT) adopts shark finning regulation (fin:carcass ratio)

2006

2007

2008

2009

EU adopts Porbeagle quota 2010

2011

The Shark Alliance Campaign is launched. Basking Shark and White Shark retention prohibited in EU Community Waters

EEA (see p.17 for country member groups established in this year)

OSPAR List of threatened and declining species includes 11 elasmobranchs

Bigeye Thresher retention prohibited (ICCAT); the EU Community Plan of Action for Sharks (CPOASharks) adopted EEA (see p.17)

Angelshark, Common Skate (see p.15), Undulate Ray, White Skate retention prohibited in all EU Community Waters; Porbeagle retention prohibited for EU vessels internationally; Spurdog 0t quota EEA (see p.17)

Common and Blackchin Guitarfishes retention prohibited in EU Community Waters contd. from p.15

2012

The General Fisheries Commission for the Mediterranean (GFCM) prohibits retention of the 24 species of elasmobranchs listed in Annex II of the Barcelona Convention SPA/BD Protocol (see Table 5)

THE EUROPEAN ELASMOBRANCH ASSOCIATION (EEA)

Established in 1996, the EEA’s objectives are to advance research, sustainable management, conservation and education of chondrichthyans throughout Europe and the Mediterranean. The EEA represents non-profit organisations, with countries represented by a single member group. There are currently 13 active member groups*: Year national member groups established –1995: 1 Deutsche Elasmobranchier-Gesellschaft (DEG)*, 2 Gruppo Ricercatori Italiani sugli Squali, Razze e Chimere (GRIS)*; 1997: 3 Shark Trust*, 4 Association Pour l’Etude et la Conservation des Sélaciens (APECs)*, 5 Associação Portuguesa para o Estudo e Conservação de Elasmobrânquios (APECE)*, 6 Shark Foundation*; 7 The Irish Elasmobranch Group (IEG)*; 2001: 8 Uno Squalo per Amico (San Marino); 2003: 9 SUBMON*; 2008: 10 Shark Lab Malta*; 2009: 11 Hai-Alliansen I Norge (HAI Norge)*; 2010: 12 Nederlandse Elasmobranchen Vereniging (NEV)*; 2016: 13 iSea*; 2019: 14 Sharks in Israel*. eulasmo.org

THREATENED ENDEMIC SKATES OF THE MEDITERRANEAN

Three species of endemic skate are found in the Mediterranean: 1 the Critically Endangered Maltese Skate Leucoraja melitensis (p.106), 2 the Endangered Rough Skate Raja radula (p.128), and 3 the Least Concern Speckled Skate Raja polystigma (p.127). While there are no specific conservation measures for the Rough Skate, the Maltese Skate is listed as a prohibited species by the GFCM.

(see above for country member groups joining in this year)

GFCM adopts FNA; Turkey prohibits retention of 14 elasmobranchs including: angelsharks (3 spp.); Giant Devil Ray; Common and Blackchin Guitarfishes; and Shortfin Mako. Whale Shark retention prohibited in European Waters

ICCAT: first International catch limit adopted in Atlantic for Blue Shark EEA (see above for country member groups joining in this year)

EU adopts Blue Shark quota

Cartilaginous Fishes Action Plan updated

IUCN Red List status

The IUCN Red List of Threatened Species is intended to be an easily and widely understood system for classifying the extinction risk of species. It divides species into nine categories:

Not Evaluated Species that have not yet been evaluated against the Red List criteria.

● Data Deficient (DD)

Species for which there is inadequate information to make an assessment of risk of extinction.

● Least Concern (LC)

Species that do not qualify for a threatened category or Near Threatened.

● Near Threatened (NT)

Species that have been evaluated as being close to, or likely to qualify for a threatened category in the near future.

● Vulnerable (VU)

Species for which the best available evidence indicates it is considered to be facing a high risk of extinction in the wild.

Northeast Atlantic

Comparison of Red List species found in both the Northeast Atlantic (drawing upon EU listings) and Mediterranean

● Endangered (EN)

Species for which the best available evidence indicates it is considered to be facing a very high risk of extinction in the wild.

● Critically Endangered (CR)

Species for which the best available evidence indicates it is considered to be facing an extremely high risk of extinction in the wild.

● Extinct in the Wild (EW)

This category is for species that are known only to survive in cultivation or captivity.

● Extinct (EX)

This category is for species when there is no reasonable doubt that the last individual has died.

For further information on all these categories and all the latest status updates visit: iucnredlist.org

Northeast Atlantic

Mention Europe in the context of shark conservation and it is inevitable that, while Europe itself extends beyond the borders of the EU, it is the EU and its fisheries interests that loom large. Widely acknowledged as a dominant force in global shark fisheries, consumption and trade, three of the 27 EU Member States rank in the top 20 shark catching nations globally, with Spain second only to Indonesia. Although the EU’s domestic target fisheries have declined along with Northeast Atlantic shark populations, the EU exerts a strong influence on global ocean policy and presents a significant voting block in environmental fora. With EU Member States keen consumers of shark products (e.g. Italy ranked 3rd for shark imports globally during 2000–2011) the significant weight of the fishing industry lobby is keenly felt.

While the EU acts to centralise many of the regulatory decisions that affect sharks, national governments have autonomy to adopt domestic legislation. Spain, for example, despite its position as a key shark catcher, has the most comprehensive domestic legislation of any EU Member State. However, Spain’s domestic Prohibition on fishing for more than 30 species is not reflected in the waters of other Northeast Atlantic countries. The result is a patchwork of measures across the region, leaving sharks to unwittingly run the gauntlet.

Between 2006–2013 the Shark Alliance campaign coordinated advocacy efforts across Europe. The CPOAShark (2009) was a key campaign objective, and the CPOA-Shark prioritised the need to tighten the EU shark finning regulation (see box, right). From 2009–2011 there was also a significant increase in the number of elasmobranchs included in the EU’s annual Fishing Opportunities (TACs and Quotas) document, listing management for Common Skate Dipturus batis (see box p.15) and Angelshark Squatina squatina for the first time. But while the CPOA had set the stage for sweeping improvements in shark fisheries management and protection policies, some 10 years after adoption, progress has generally been disappointing and many elasmobranchs remain without effective management.

Beyond a plethora of specific EU instruments, the waters of the Northeast Atlantic fall under the jurisdiction of the OSPAR and HELCOM Regional Seas Conventions and multiple Regional Fishery Bodies, and within the remit of intergovernmental organisations (see Table 6, p.26). Each of these has the capacity to adopt measures which may be binding or non-binding. These additional instruments also address the fact that not all the waters of the Northeast Atlantic are under the jurisdiction of EU Member States. For example, the Exclusive Economic Zone (EEZ) of Morocco, a country with significant shark

fishing interests, nudges into the south of the region. The longline fleets of southern Europe and Morocco dominate the shark landings of the North Atlantic. Acknowledging this, in recent years advocacy efforts have focused on the adoption of binding ICCAT Recommendations for Blue Shark Prionace glauca and Shortfin Mako Isurus oxyrinchus. In 2019 a significant milestone was reached with the adoption of the first international catch limits for Blue Sharks, while Shortfin Makos remain without catch limits despite scientific advice for a prohibition in the North Atlantic.

SHARK FINNING

Shark finning is the practice of removing a shark’s fins at sea and discarding the carcass overboard. It is a highly wasteful practice and a major barrier to effective fisheries management (Fin Guide p.369).

In 2003 the EU adopted a Shark Finning Regulation which, while banning finning itself, permitted removal of fins at sea, on the proviso that both fins and carcass were retained. Compliance was monitored through a fin : carcass weight ratio, however this was set higher (5% whole weight) than other fishing nations (e.g. US 5% dressed weight). In addition, the EU permitted fins and carcasses to be landed in separate ports – further complicating the enforcement process and allowing less scrupulous operators to land fins from finned and discarded sharks.

Fins Naturally Attached (FNA) is now widely accepted as best practice for regulating shark finning as it eases enforcement, aids species identification, prevents high grading (the mixing of fins from high value species with smaller, or lower value carcasses), which in turn improves landings data. Following an effective campaign by the Shark Alliance, FNA was adopted by the EU in 2013 and is applicable to the EU fleet globally. FNA has also been adopted by NEAFC (2015), and GFCM (2018).

Partial cut – fins are cut along the majority

The Mediterranean

Where the Northeast Atlantic has a relatively clear focal point in the form of the EU, the Mediterranean is the counterpoint: a highly complex, multijurisdictional region surrounded by over 20 countries and territories, across three continents. European influences are dominant in the west, with the waters and fisheries of the 11 EU Mediterranean coastal states addressed in the main through EU regulations. Non-EU countries are governed by domestic legislation and commitments to regional bodies and Multilateral Environmental Agreements (MEAs). Many Mediterranean countries are Party to ICCAT, all are Party to the Barcelona Convention, GFCM and its elasmobranch measures, and all bar Turkey are Signatories to the Convention on Migratory Species (CMS).

There has been a significant decline in species richness throughout the Mediterranean, coinciding with reported declines in elasmobranch abundance. More than a decade since Mediterranean chondrichthyans were first assessed, the IUCN reports that there is no sign of improvement in the regional status of sharks, skates and rays. Overfishing is the key threat, complicated by the fishing sector’s historically important economic role in the region.

Subsistence or small-scale fisheries (SSF) are deeply rooted in the fabric of the Mediterranean, with SSF playing a significant social and economic role. Indeed, 84% of the fishing fleet, some 70,000 vessels, operate in SSF, generally supplying local markets. Perhaps the key constraint to delivering effective conservation or management activities is reaching this SSF sector, with fishing villages often located along remote coastlines. The dispersed nature of the region’s fishing activity makes quantifying species-specific elasmobranch bycatch challenging. This is further hampered by the significant level of aggregated catch reporting: over 65% of all reported elasmobranch catches in the Mediterranean are landed in the aggregate.

Because a number of governments have lagged behind in implementing existing regulations (primarily GFCM/42/2018/2), fishers and fisheries enforcement officers may be unaware of existing regional recommendations –the result often being the sale of all bycatch regardless of regional conservation management status. Efforts to address the commercialization of threatened species have often been led by the NGO community and have included: an increase in bycatch observer coverage; education programmes for fishers and retailers; and an expansion of advisory and identification materials (see Table 6).

While overfishing remains the primary threat, habitat degradation is also a key concern. The Barcelona Convention’s Specially Protected Areas Regional Advisory Centre (RAC/ SPA) provides a valuable coordinating role in the region.

GUITARFISHES

Giant Guitarfishes are the world’s most threatened marine fishes, and the Critically Endangered Blackchin Guitarfish Glaucostegus cemiculus (p.181) is native to the Mediterranean. Also found in region are the Endangered Common Guitarfish Rhinobatos rhinobatos (p.182). Both species are caught and retained in mixed and target fisheries along the North African coast. Guitarfishes are a high value ray with the meat consumed locally. Little is known about actual landings, catch composition, seasonality, market demand, or trade.

These guitarfishes are listed as prohibited species by the GFCM, but this regulation is poorly implemented, leaving them exposed to unregulated fishing pressure.

GIANT DEVIL RAY

Capture and retention of 30 Giant Devil Ray Mobula mobular (p.163) by a Turkish vessel in 2017 exposed troubling gaps in the compliance with GFCM regulations, which prohibit their landing and selling. Amongst the world’s least fecund marine fish, the Giant Devil Ray is vulnerable to overexploitation due to its late maturity, lengthy gestation, and exceptionally small numbers of offspring – typically producing one pup every two to three years. Landing and selling the Giant Devil Ray is banned in the Mediterranean under GFCM/42/2018/2, yet bycatch still occurs. Triggered by the 2017 Turkish landings, and associated conservation concerns, the Turkish Government added the Devil Ray to their domestic prohibited species list in 2018.

Securing conservation management

Safeguarding the future of elamobranchs is a long-term commitment, particularly when considering waters as depleted as the Mediterranean, and as pressured as the Northeast Atlantic. Current progress reflects decades of effort, yet many species and their critical habitats remain vulnerable in the absence of management. It is clear that existing conservation measures are not enough.

Conservation communities

As shark populations have declined, there has been a growth in the number of groups and organisations voicing interest in elasmobranchs, including environmental Non-Governmental Organisations (NGOs), the research community, fisheries interest groups, governments and an increasingly vocal general public.

The public attitude to sharks can be as diverse as ‘the public’ themselves which, in the context of shark conservation, might encompass a range of constituencies from SCUBA divers and recreational sea anglers, to wildlife enthusiasts, beach-goers and the consumer.

Over the past 25 years, the public’s attitude to the natural environment has evolved and public influence on decision-making with it. Previously, sharks were most likely encountered on a visit to an aquarium, as a menu item, or on the end of a fishing line. This hasn’t much changed, although now sharks are also a big ticket for ecotourism – with operators in the region responding to growing interest in seeing sharks in the wild.

This evolving fascination with sharks is most clearly reflected in the proliferation of citizen science projects. These are often sightings projects which engage ‘citizens’ on land and sea as recorders, noting occasional sightings (see Box) or undertaking regular surveys. The result is many more eyes on the water, strandline or fish market, and collation of a greater volume of data and information over a wider geographic range than could be achieved by the research community working alone. The widescale use of smart phones adds valuable technical capacity, with cameras, GPS and access to multiple recording platforms and public fora providing new and growing data streams. Similarly, social media feeds represent a surprising source of records and sightings.

Yet, despite a more informed public, the consumer market for shark products remains strong – with shark meat still popular and shark products still widely used. At the same time, sectors of the recreational community continue to prize shark angling, with the targeting of vulnerable species, such as Porbeagle Lamna nasus, evident in the Northeast Atlantic.

Securing change

Public campaigning is often a key element of conservation change. Indeed, effective campaigning is a common theme underpinning many of the ‘wins’ illustrated on the earlier timeline. Mobilisation of shark conservation groups across Europe under the Shark Alliance campaign is a positive example. But the conservation landscape is changing. Where the shark fin trade was the key driver, retention of sharks for meat is now a major consideration. In this context the conservation community must update its narrative and avoid oversimplification of messaging. There is not a single solution to the challenges of shark conservation, and campaigns must be nuanced to reflect this. Simply calling for bans, whether on specific products or the shark fin trade, will not stop sharks being caught or killed. Such a limited approach may, in fact, inadvertently divert attention from the real threat: that despite some progress, most shark, skate and ray populations continue to be fished with no limits.

CITIZEN SCIENCE: THE GREAT EGGCASE HUNT

The Shark Trust’s public recording project, the Great Eggcase Hunt, gathers georeferenced data for shark, skate and chimaera eggcases found primarily around the United Kingdom, but increasingly from across Europe, the Mediterranean, and beyond. While acting as a useful tool to engage the public in shark conservation, records can also indicate presence and diversity of oviparous species in different regions.

Eggcases (or ‘mermaid’s purses’) vary according to species (see p.42). By examining various features (size, shape, whether they have horns or curled tendrils, presence of lateral keels etc.) it is possible to determine which species the eggcase came from. To assist with identification, the Shark Trust has developed resources for those eggcases most likely to be found on northwest European shores. Once identified, submission to the Great Eggcase Hunt database is encouraged, with a photo to allow for reliable verification of records. www.eggcase.org

Challenging times

Securing effective conservation management and seeking implementation of and compliance with existing measures at national, regional and international levels is crucial. Collaborative conservation strategies, such as that published by the Global Sharks and Rays Initiative, and species-specific regional action plans (see angelshark box below) provide structured roadmaps for delivery of conservation objectives. The findings of such reports, where effectively communicated, can help secure much needed policy decisions.

Constraints to effective conservation management are commonplace, but may also differ between the Northeast Atlantic, particularly EU waters, and those of the Mediterranean. These constraints reflect political, economic, legal and social factors. In a region as diverse as the Mediterranean, cultural and linguistic diversity are also key considerations; efforts must be taken to prevent language from acting as a barrier to delivering effective conservation management (for example, by translating identification guides and advisory materials into multiple languages). Efforts to secure and maintain the focus of decision-makers, and to ensure that governments engage across their institutions – linking the decisions of Environment Ministries with delivery through Fisheries Ministries – is key. The implementation of CITES listings for elasmobranchs – such as controls on the international trade in Appendix II listed species – is an example of this. Furthermore, where CITES-listed species are caught and landed from a country’s EEZ but not traded internationally, there is an even greater urgency to see domestic management enforced. The Blackchin Guitarfish Glaucostegus cemiculus (CITES Appendix II) and Common Guitarfish Rhinobatos rhinobatos provide a example of this disconnect: both species are landed in significant numbers on the North Africa coast, despite being prohibited through regional regulations adopted by the GFCM Parties.

This is where conservation advocates can come to the fore, moving swiftly across the science-policy interface, holding governments and the fishing industry to account for commitments made, while championing scientific advice as the foundation of effective conservation and fisheries management.

Finally…

The past 25 years have seen significant change in elasmobranch conservation and management, and shifts in government positions and public perceptions. Advocacy and ‘awareness’ campaigns have established an understanding that chondrichthyans are vital to a healthy marine environment.

Yet, while we continue to focus on much-needed catch limits, we must also ensure governments and regulators demand improved landings data, and end the reporting of landings using aggregated categories. Better data can underpin more accurate stock assessments – the building blocks of clear, unambiguous scientific advice. We need governments to adhere to the science, as it provides the basis for sustainable fisheries and alerts to the need for strict protection for more vulnerable species. Maintaining the economic viability of a fishery should not be achieved by continued retention of threatened species.

With many elasmobranchs caught as incidental bycatch, bolstering catch limits with bycatch mitigation and avoidance measures is key. This may include the adoption of spatial, temporal and technical measures, within EEZs and on the high seas.

Conservation and management of chondrichthyans is a complex business. But it is relatively simple to get involved, whether you are a researcher, an advocate, a conservationist or simply passionate about sharks and their relatives. The key thing is to engage.

COLLABORATION: ANGELSHARKS

Angelsharks have been identified as one of the most threatened families of chondrichthyans in the world, with many species requiring urgent conservation action. The Angel Shark Conservation Network (ASCN) was established to facilitate dialogue and information sharing on angelshark conservation efforts, particularly throughout the range of the three Critically Endangered species found in the Eastern Atlantic and Mediterranean Sea: Squatina aculeata, S. oculata and S. squatina.

The Mediterranean Angel Sharks: Regional Action Plan sets out a roadmap focusing efforts to restore these enigmatic species to robust populations in the region. Goals are centred around the predominant threats to angelsharks: fisheries and habitat degradation, with an underlying goal for the implementation of legislation and regulations to ensure both the species and their critical habitats are better protected. To allow a tailored approach in this multijurisdictional region and engage regional stakeholders, a series of Subregional Action Plans focus on subareas of the Mediterranean which are considered high priority for conservation action. www.sharktrust.org/angelsharks

Conservation and fisheries management resources

Sharpnose Sevengill Shark

Heptranchias perlo

Spurdog Squalus acanthias

Gulper Shark

Leafscale Gulper Shark

Portuguese Dogfish

Centrophorus granulosus

Centrophorus squamosus

Centroscymnus coelolepis

Angular Roughshark Oxynotus centrina

Sawback Angelshark Squatina aculeata

Smoothback Angelshark Squatina oculata

Angelshark Squatina squatina

Whale Shark Rhincodon typus

Smalltooth Sandtiger Odontaspis ferox

Bigeye Thresher Alopias superciliosus

Common Thresher Alopias vulpinus

Basking Shark Cetorhinus maximus

White Shark Carcharodon carcharias

Shortfin Mako Isurus oxyrinchus

Longfin Mako Isurus paucus

Porbeagle Lamna nasus

Tope Galeorhinus galeus

Starry Smoothhound Mustelus asterias

Smoothhound Mustelus mustelus

Blackspotted Smoothhound Mustelus punctulatus

Silky Shark Carcharhinus falciformis

Oceanic Whitetip Shark Carcharhinus longimanus

Dusky Shark Carcharhinus obscurus

Sandbar Shark Carcharhinus plumbeus

Blue Shark Prionace glauca

Scalloped Hammerhead Sphyrna lewini

Great Hammerhead Sphyrna mokarran

Smooth Hammerhead Sphyrna zygaena

Common Skate complex: Blue Skate Flapper Skate

Dipturus batis Dipturus intermedius

Black Skate Dipturus nidarosiensis

Maltese Skate

Sandy Skate

Leucoraja melitensis

Leucoraja circularis

Thornback Skate Raja clavata

Spotted Skate Raja montagui

White Skate Rostroraja alba

Spiny Butterfly Ray Gymnura altavela

Giant Manta Ray Mobula birostris

Giant Devil Ray Mobula mobular

Smalltooth Sawfish Pristis pectinata

Largetooth Sawfish Pristis pristis

Blackchin Guitarfish

Glaucostegus cemiculus

Common Guitarfish Rhinobatos rhinobatos

App

App

Annex II*

Key wildlife treaties and management instruments (Table 3) list a fraction of the species found in the region, and offer no protection to many of the most vulnerable. As our knowledge on elasmobranch populations and the threats they face expands, and IUCN Red List reviews are undertaken, it becomes clear that many species currently listed as Data Deficient are likely threatened and the scope of conservation actions should be expanded. Listing species on a regulation, however, does not guarantee implementation.

Table 5. Key wildlife treaties and management instruments

The Convention on International Trade in Endangered Species of Wild Fauna & Flora (CITES)

CITES Appendix I: International commercial trade is prohibited except in exceptional circumstances.

CITES Appendix II: Trade is controlled under specific conditions using permits.

The Convention on the Conservation of Migratory Species of Wild Animals (CMS)

CMS Appendix I: Species are to be strictly protected.

CMS Appendix II: CMS Parties have committed to work internationally towards conservation.

The Memorandum of Understanding on the Conservation of Migratory Sharks (Sharks MOU)

CMS MoU Annex I: lists species that have an unfavorable conservation status and which require international agreements for their conservation and management, or would significantly benefit from the international cooperation that could be achieved by an international agreement.

Barcelona Convention for the Protection of the Mediterranean Protocol concerning Specially Protected Areas and Biological Diversity (SPA/BD)

Annex II: Endangered and threatened species

Annex III: Species whose exploitation is regulated

OSPAR

The OSPAR list of threatened and/or declining species.

International Commission for the Conservation of Atlantic Tunas (ICCAT)

ICCAT: o management measure applies; ** exemptions apply; x prohibitions apply. Shark finning is regulated in the ICCAT Convention Area through a fin to carcass ratio.

General Fisheries Commission for the Mediterranean (GFCM)

Adopted a prohibition on the retention, transshipment, landing, storage, display and sale of 23 species of elasmobranchs listed on Barcelona SPA/BD Annex II through GFCM/36/2012/3 (since superseded by GFCM/42/2018/2).

Fins Naturally Attached was adopted in 2018.

EU Regulations

Total allowable catches (TACs) or fishing opportunities, are catch limits (expressed in tonnes) that are set for most commercial fish stocks, but many elasmobranchs have no catch limits, and exceptions may apply. TACs and Quotas are subject to annual/biennial change. Details on current TACs and Quotas can be found on eur­lex.europa.eu and condensed versions are presented as Fisheries Advisories (Table 6, p.26).

Prohibited Species are listed in four key Council Regulations: (EU) No. 2015/812: EU Landings Obligation (EU) No. 2019/1241: EU Technical Measure (EU) No. 2015/2102: EU Regulation on GFCM (EU) 2018/2025: Fishing opportunities for Union fishing vessels for certain deepsea fish stocks including sharks.

EU Landings Obligation (EU) No 1380/2013: The landing obligation requires all catches of regulated commercial species onboard to be landed and counted against quota, unless the species is prohibited or exemptions apply.

This table does not present national conservation or management legislation but signposts international and regional instruments, resources and recording platforms. All listings are regularly subject to change.

Table 6. International and regional instruments, resources and recording platforms

Organisation Further information

Fisheries Management

Regional Fishery Bodies:

Website

GFCM General Fisheries Commission for the Mediterranean fao.org/gfcm

ICCAT International Commission for the Conservation of Atlantic Tunas iccat.int

ICES International Council for the Exploration of the Sea ices.dk

LDAC Long Distance Advisory Committee ldac.eu

NEAFC North East Atlantic Fisheries Committee neacf.org

NWWAC North Western Waters Advisory Committee nwwac.org

EU Regulations:

EU Deepsea fish stocks Regulation (EU) 2018/2025

EU Landings Obligation Regulation (EU) No. 2015/812

EU Regulation on GFCM Regulation (EU) No. 2015/2102

EU Technical Measure Regulation (EU) No. 2019/1241

Fishery Advisories

EU Regulations and handling guides. Updated annually, multilingual sharktrust.org/advisories

FAO FishStat Plus Fishery statistical time series fao.org/fishery/statistics

TACs and Quotas Council Regulation (EU) No. 2020/123 NB: updated annually eur­lex.europa.eu

Multilateral Environment Agreements

Barcelona

The Convention for the Protection of the Marine Environment and the Coastal Region of the Mediterranean, Mediterranean Action Plan The Regional Activity Centre for Specially Protected Areas (RAC/SPA)

unenvironment.org/unepmap rac­spa.org

Bern The Convention on the Conservation of European Wildlife and Natural Habitats coe.int/en/web/bern­convention

CBD Convention on Biological Diversity cbd.int

CITES The Convention on International Trade in Endangered Species of Wild Fauna and Flora cites.org

CMS The Convention on the Conservation of Migratory Species of Wild Animals cms.int

CMS MoU The Memorandum of Understanding on the Conservation of Migratory Sharks cms.int/sharks

HELCOM

The Helsinki Convention on the Protection of the Marine Environment of the Baltic Sea Area helcom.fi

OSPAR Convention for the Protection of the Marine Environment of the Northeast Atlantic ospar.org

UNEP/RSCAPs United Nations Environment Programme/Regional Seas Programme unenvironment.org/exploretopics/oceans­seas

Resources and contacts

ASCN Angel Shark Conservation Network angelsharknetwork.com/#maps

EEA European Elasmobranch Association eulasmo.org

EUAC European Union of Aquarium Curators euac.org

GSRI Global Sharks and Rays Initiative globalsharksraysinitiative.org

Organisation Further information Website

Resources and contacts (contd.)

IUCN Red List of Threatened Species For species’ conservation status iucnredlist.org

IUCN Shark Specialist Group For a wealth of information on biology, distribution, population status and conservation actions iucnssg.org

Shark League Shark League for the Atlantic and Mediterranean sharkleague.org

Shark Trust Science-based, shark conservation charity founded in 1997 sharktrust.org

TRAFFIC Wildlife Trade Specialists traffic.org

Recording platforms

ASCN Angel Shark Sightings Map sites.zsl.org/angelsharks

CIESM The Mediterranean Science Commission’s Shark Task Force records observations of rare species ciesm.org/forums/Sharks.html

MECO The Mediterranean Elasmobranch Citizen Observation project facebook.com/theMECOproject

MEDLEM Mediterranean Large Elasmobranch Monitoring arpat.toscana.it/medlem

Shark Trust Elasmobranch Public Sightings Database sharktrust.org/sightings­database

The Great Eggcase Hunt A Shark Trust initiative: identification and recording resources (see p.22) eggcase.org

Further reading

Given the plethora of papers published on chondrichthyan issues in the Northeast Atlantic and Mediterranean, no attempt has been made to present a comprehensive reference list – and no offence is intended to any authors omitted. This list is a selection of publications which, in the main, address more wide reaching issues such as regional population status, actions plans or trade data.

Barker, J. et al. 2016. Angelshark Action Plan for the Canary Islands. Zoological Society of London, United Kingdom.

Bradai M.N., et al. 2012. Elasmobranch of the Mediterranean and Black sea: status, ecology and biology. Bibliographic analysis. Studies and Reviews.

General Fisheries Commission for the Mediterranean. No. 91. Rome, FAO. 103pp.

Bräutigam, A. et al. 2015. Global Priorities for Conserving Sharks and Rays: a 2015–2025 strategy. The Global Sharks and Rays Initiative. 28pp.

Cavanagh, R.D. & Gibson, C. 2007. Overview of the Conservation Status of Cartilaginous Fishes (Chondrichthyans) in the Mediterranean Sea. IUCN, Gland, Switzerland and Malaga, Spain. Vi + 42pp.

Dent, F. & Clarke, S. 2015. State of the global market for shark products. FAO Fisheries and Aquaculture Technical Paper No. 590. Rome, FAO. 187pp.

Dulvy, N.K. et al. 2014. Extinction risk and conservation of the world’s sharks and rays. eLife, 3: e00590.

Dulvy N.K. et al. 2016. The Conservation Status of Sharks, Rays and Chimaeras in the Mediterranean Sea [Brochure]. Malaga, Spain: IUCN.

FAO. 2018. The State of Mediterranean and Black Sea Fisheries. General Fisheries Commission for the Mediterranean. Rome. 172pp. Licence: CC BY-NCSA 3.0 IGO.

FAO FishStat Plus – Universal software for fishery statistical time series. Rome. www.fao.org/fishery/statistics/software/fishstat/en (updated to 2017).

Ferretti, F. et al. 2016. Squatina oculata The IUCN Red List of Threatened Species 2016: e.T61418A16570000. Mediterranean assessment.

Gibson, C., Valenti, S.V., Fordham, S.V. and Fowler, S.L. 2008. The Conservation of Northeast Atlantic Chondrichthyans: Report of the IUCN Shark Specialist Group Northeast Atlantic Red List Workshop. IUCN Species Survival Commission Shark Specialist Group. Newbury, UK. viii + 76pp.

Gordon, C.A. et al. 2017. Eastern Atlantic and Mediterranean Angel Shark Conservation Strategy. The Shark Trust, United Kingdom.

Gordon, C.A. et al. 2019. Mediterranean Angel Sharks: Regional Action Plan The Shark Trust, United Kingdom. 36pp.

IUCN/SSC. 2008. Strategic Planning for Species Conservation: A Handbook. Version 1.0. Gland, Switzerland: IUCN Species Survival Commission. 104pp.

IUCN 2020. The IUCN Red List of Threatened Species. Version 2020–2. https://www.iucnredlist.org.

Lawson, J.M. & Fordham, S.V. 2018. Sharks Ahead: Realizing the Potential of the Convention on Migratory Species to Conserve Elasmobranchs. Shark Advocates International, The Ocean Foundation, Washington, DC, USA. 76pp.

Mancusi, C. et al. 2020. MEDLEM database, a data collection on large elasmobranchs in the Mediterranean basin. Mediterranean Marine Science 21(2), 276-288.

Nieto, A. et al. 2015. European Red List of marine fishe s. Luxembourg: Publications Office of the European Union.

Okes, N. & Sant, G. 2019. An Overview of Major Shark Traders, Catchers and Species. TRAFFIC, United Kingdom.

SPA/RAC–UN Environment/MAP. 2012. Guidelines for shark and ray recreational fishing in the Mediterranean

SPA/RAC–UN Environment/MAP. 2020. Action Plan for the Conservation of Cartilaginous Fishes (chondrichtyans) in the Mediterranean Sea; by: Bradai, M N., Ed SPA/RAC. Tunis, 18 pp.

Turkoglu, M. et al. (eds.) 2018. Marine Ecology: Biotic and Abiotic Interactions. IntechOpen.

HOW TO USE THIS BOOK

The structure of this book will be familiar to anyone who has used other species field guides. Where this field guide differs from most, is that the illustrated dichotomous (branching) key guides are found throughout the book. The first key (pp.43–44) starts with the cartilaginous fish, which takes you to further key guides, first to orders, then families and genera and finally species. We recommend always using the dichotomous key when trying to identify a particular species (even if you already think that you know what it is).

To further help identification, spreads throughout the guide show similar looking species together for comparison.

Each order and family has its own introduction with the overall habitat and ecology for the group as well as key identification characters. This introductory section also reviews briefly the biology and status of the taxonomic group.

The introduction to each family is followed by a page entry for every species occurring in the region. The species is identified by one widely used English name, its scientific name and its FAO 3-Alpha code. Species appear in alphabetical order of their scientific names within each family, firstly the genus, then the species name.

Each species description is accompanied by colour illustrations of the lateral and ventral view for sharks (except for angelsharks which have dorsal and ventral views), a lateral view for chimaeras, and dorsal and ventral views for batoids. Scale bars indicate the approximate sizes of the species (in cm) at different ages, when known. Total body length (TL) measurements are given in cm, measured as the ‘point to point’ distance (i.e. not over the curve of the body) from the tip of the snout to the tip of the caudal fin (tail). Where available, these include total length at birth, maturity and maximum size. For chimaeras, body length (BDL) and precaudal length (PCL) are sometimes used (see p.363). For devil rays, eagle rays, cownose rays and stingrays, disc width (DW) is used as the overall measurement. For egg-laying species, we have tried to indicate the size of eggcase and length of young when they hatch and there are separate sections comparing these eggcases (see later in this introduction). ‘Mature’ indicates the size at which the species first reaches sexual maturity (this is often different for males and for females; the latter are generally larger) and where possible we have indicated for both. The ‘Maximum’ length is often the largest recorded, not the maximum that a species could reach (which may be much larger). Weights have not been given as they are not often

recorded in scientific literature because they are highly variable, depending upon the time of year, state of pregnancy, recent meals and so on. Length, on the other hand, is a more constant, reliable and useful measure of size (and age) of animal. Length:weight conversion tables are available for many species, to allow anglers to estimate the weight of their catch without removing it from the water and possibly injuring the animal.

The description text includes each species’ shape and other distinctive features. The drawings on pp.30–34 identify the body parts referred to in the text. There are also sections on colour and dentition. Dentition descriptions include tooth counts (where known); this is the total number of tooth rows along both jaw lines (left and right) of the upper and lower jaws. This figure is not the total number of teeth to be found in the jaw as this would include all the series of teeth in the rows, it is only the outermost series, the functional teeth, that are the rows counted (see p.35). Throughout the book, key guides help identify teeth for a particular group or, in the case of sharks, to species level. These key guide dentition illustrations are of all the functional teeth, from one side of the upper and lower jaws; accompanying scale bars indicate the size of these adult teeth.

Distribution is provided as a map. It is quite possible that many species have a wider regional distribution than illustrated, especially in poorly recorded species. Solid blue in the maps refers to known distribution, lighter blue is possible distribution.

Distribution is indicated in dark blue for known occurrence and light blue for possible distribution. The Azores distribution is sometimes shown as an inset, as above, where there is a wideranging east–west distribution.

Habitat describes where the animal most commonly lives, including the depths below sealevel, in metres, where it has been reported.

Biology summarises information known about species reproduction; behaviour, including movements and migrations; and characteristic feeding activities and prey. Also included in this section is information on eggcases, where applicable. Pages throughout the book illustrate these eggcases, where they are known, drawn to scale for that page. For more information on eggcases and to record any eggcase found, visit www.eggcase.org (see p.22).

IUCN Red List status (see also pp.24–25) is current in July 2020, but updated twice a year (see www.redlist.org for more recent assessments). Brief information about the conservation, management and fisheries status of the animal is also given.

On the right hand side of the page there is a summarised visual guide in the form of icons Starting from the top is the ease of identification icon, then the depth guide which shows the range of depth recorded for that species, and sometimes at which depths it more commonly occurs (darker yellow tint). Below that are a series of icons (see right) for temperature range, habitat, reproductive method, litter/eggcase number (if known), maturity age, maximum age, where the species is found in the water column and finally its diet. NR is used to denote data not recorded. The last set of icons relate to its conservation and management status.

The glossary of technical terms is provided on pp.352–356. Regional maps showing sea depths, sea currents, sea salinity, average sea temperatures and marine climate zones are on pp.357–359. To help with recording, the field observation section covers chimaeras, batoids and sharks, pp.361–370, and finally a section on shark fins on pp.371–378.

EASE OF IDENTIFICATION

Should be easy to identify

Often a close inspection needed to identify

Difficult to identify

Temperature range

Habitat

Mud / Sand / Rock / Pelagic / Kelp forests

Reproductive method

Number of young/eggcases

Oviparous / Oviviparous / Viviparous /

Oophagus

Gestation period (months)

Maturity age

Maximum age

Where found in water column

Surface (0–200m) / midwater (>200m) / bottom (lives on bottom)

Diet

Shellfish / scavenger / piscivorous teleosts / crustaceans / invertebrates (general) / cephalopods / mammals-birds / piscivorous chondrichthyans / parasitic / planktivorous

IUCN RED LIST STATUS

(see definitions on p.18)

Not Evaluated

Data Deficient

Least Concern

Near Threatened

CITES APPENDIX LISTING

CITES Appendix I International commercial trade is prohibited.

CITES Appendix II Trade is regulated through permits.

CMS APPENDIX LISTING

Appendix I Threatened migratory species

Appendix II Migratory species whose status would benefit from international cooperation

BARCELONA CONVENTION LISTING

Annex II Endangered and threatened species

Annex III Species whose exploitation is regulated

Topography

bulbous tip sharp denticles

Front tenaculum

position of front tenaculum

eye

snout

vent fin spine

first dorsal fin

second dorsal fin

lateral line canal

mouth

gill opening soft operculum

prepelvic tenaculum

clasper

anal fin pelvic fin

Ventral view of claspers

Lateral view – chimaera

pectoral fin caudal filament

Prepelvic tenaculum

upper lobe of caudal fin

lower lobe of caudal fin

nostril lateral line canals

tongue

vomerine dental plate palatine dental plate

Ventral view – chimaera mouth

mandibular dental plate

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